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Floristic Evidence for Alternative Biome States in Tropical Africa

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Floristic Evidence for Alternative Biome States in Tropical Africa Floristic evidence for alternative biome states in tropical Africa J. C. Alemana,b, A. Fayollea,1, C. Favierc, A. C. Staverd, K. G. Dextere,f, C. M. Ryane, A. F. Azihoug, D. Baumanh,i, M. te Beestj,k,l, E. N. Chidumayom, J. A. Comiskeyn,o, J. P. G. M. Cromsigtj,k,p, H. Dessardq,r, J.-L. Douceta, M. Finckhs, J.-F. Gilleta, S. Gourlet-Fleuryq,r, G. P. Hempsont, R. M. Holdou, B. Kirundav, F. N. Kouamew, G. Mahya, F. Maiato P. Gonçalvesx, I. McNicole, P. Nieto Quintanoe, A. J. Plumptrev,y,z, R. C. Pritcharde,aa, R. Revermanns,bb, C. B. Schmittcc,dd, A. M. Swemmeree, H. Talilaff, E. Woollene, and M. D. Swainegg aGembloux Agro-Bio Tech, Université de Liège, 5030 Gembloux, Belgium; bDépartement de Géographie, Université de Montréal, Montréal, QC H2V 0B3, Canada; cInstitut des Sciences de l’Evolution–Montpellier, CNRS, Ecole Pratique des Hautes Etudes, Institut de Recherche pour le Développement, Université de Montpellier, 34000 Montpellier, France; dDepartment of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06520; eSchool of GeoSciences, University of Edinburgh, EH8 9YL Edinburgh, United Kingdom; fTropical Diversity Section, Royal Botanic Garden Edinburgh, EH3 5LR Edinburgh, United Kingdom; gLaboratory of Applied Ecology, Faculty of Agronomic Sciences, University of Abomey-Calavi, 01 BP 526 Cotonou, Benin; hEnvironmental Change Institute, School of Geography and the Environment, University of Oxford, OX1 3QY Oxford, United Kingdom; iPlant Ecology and Biogeochemistry, Université Libre de Bruxelles, 1050 Bruxelles, Belgium; jCopernicus Institute of Sustainable Development, Utrecht University, 3584 CS Utrecht, The Netherlands; kCentre for African Conservation Ecology, Nelson Mandela University, 6031 Port Elizabeth, South Africa; lGrasslands-Forests-Wetlands Node, South African Environmental Observation Network, 3201 Pietermaritzburg, South Africa; mMakeni Savanna Research Project, Ridgeway, 1001 Lusaka, Zambia; nInventory and Monitoring Program, National Park Service, Fredericksburg, VA 22405; oSmithsonian Institution, Washington, DC 20002; pDepartment of Wildlife, Fish, and Environmental Studies, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden; qForêts et Sociétés, Centre de Coopération Internationale en Recherche Agronomique Pour le Développement, Université de Montpellier, 34000 Montpellier, France; rCentre de Coopération Internationale en Recherche Agronomique Pour le Développement, Forêts et Sociétés, 34398 Montpellier, France; sBiodiversity, Evolution and Ecology of Plants, Institute of Plant Science and Microbiology, University of Hamburg, 22609 Hamburg, Germany; tCentre for African Ecology, School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, 2000 Johannesburg, South Africa; uOdum School of Ecology, University of Georgia, Athens, GA 30602; vWildlife Conservation Society, Kampala, Uganda; wNature Sciences Unit, University Nangui Abrogoua, 31 BP 165 Abidjan, Côte d’Ivoire; xHerbário do Lubango, Instituto Superior de Ciências da Educação da Huíla, C.P. 230 Lubango, Angola; yHead Key Biodiversity Area Secretariat, c/c BirdLife International, CB2 3QZ Cambridge, United Kingdom; zConservation Science Group, Zoology Department, Cambridge University, CB2 3EJ Cambridge, United Kingdom; aaGlobal Development Institute, University of Manchester, M13 9PL Manchester, United bb cc Kingdom; Faculty of Natural Resources and Spatial Sciences, Namibia University of Science and Technology, Windhoek, Namibia; Center for ENVIRONMENTAL SCIENCES Development Research, University of Bonn, 53113 Bonn, Germany; ddNature Conservation and Landscape Ecology, University of Freiburg, 79106 Freiburg, Germany; eeSouth African Environmental Observation Network, SAEON Ndlovu Node, Phalaborwa, 1390, South Africa; ffDepartment of Ecotourism and Biodiversity Conservation, College of Agriculture and Natural Resource, Madda Walabu University, Bale Robe, Ethiopia; and ggInstitute of Biological and Environmental Sciences, University of Aberdeen, AB24 3FX Aberdeen, United Kingdom Edited by Robert John Scholes, University of the Witwatersrand, Wits, South Africa, and approved September 7, 2020 (received for review June 8, 2020) The idea that tropical forest and savanna are alternative states is crucial to how we manage these biomes and predict their future Significance under global change. Large-scale empirical evidence for alterna- tive stable states is limited, however, and comes mostly from We develop a biogeographic approach to analyzing the pres- the multimodal distribution of structural aspects of vegetation. ence of alternative stable states in tropical biomes. Whilst These approaches have been criticized, as structure alone cannot forest–savanna bistability has been widely hypothesized and separate out wetter savannas from drier forests for example, and modeled, empirical evidence has remained scarce and contro- there are also technical challenges to mapping vegetation structure versial, and here, applying our method to Africa, we provide in unbiased ways. Here, we develop an alternative approach to de- large-scale evidence that there are alternative states in tree limit the climatic envelope of the two biomes in Africa using tree species composition of tropical vegetation. Furthermore, our species lists gathered for a large number of forest and savanna sites results have produced more accurate maps of the forest and distributed across the continent. Our analyses confirm extensive cli- savanna distributions in Africa, which take into account dif- matic overlap of forest and savanna, supporting the alternative sta- ferences in tree species composition, and a complex suite of ble states hypothesis for Africa, and this result is corroborated by determinants. This result is not only important for under- paleoecological evidence. Further, we find the two biomes to have standing the biogeography of the continent but also, to guide highly divergent tree species compositions and to represent alterna- large-scaled tree planting and restoration efforts planned for tive compositional states. This allowed us to classify tree species as the region. forest vs. savanna specialists, with some generalist species that span both biomes. In conjunction with georeferenced herbarium records, Author contributions: J.C.A., A.F., and M.D.S. designed research; J.C.A., A.F., A.F.A., we mapped the forest and savanna distributions across Africa and D.B., M.t.B., E.N.C., J.A.C., J.P.G.M.C., H.D., J.-L.D., M.F., J.-F.G., S.G.-F., G.P.H., R.M.H., quantified their environmental limits, which are primarily related to B.K., F.N.K., G.M., F.M.P.G., I.M., P.N.Q., A.J.P., R.C.P., R.R., C.B.S., A.M.S., H.T., E.W., and M.D.S. performed research; J.C.A., A.F., and C.F. contributed new reagents/ precipitation and seasonality, with a secondary contribution of fire. analytic tools; J.C.A. and A.F. analyzed data; A.C.S., K.G.D., C.M.R., and D.B. com- These results are important for the ongoing efforts to restore African mented on the paper; A.F.A., M.t.B., E.N.C., J.A.C., H.D., J.-L.D., J.-F.G., B.K., F.N.K., ecosystems, which depend on accurate biome maps to set appropri- G.M., F.M.P.G., I.M., P.N.Q., R.C.P., R.R., H.T., E.W., and M.D.S. contributed to data; ate targets for the restored states but also provide empirical evidence J.P.G.M.C., M.F., S.G.-F., G.P.H., R.M.H., A.J.P., C.B.S., and A.M.S. contributed to data and commented on the paper; and J.C.A., A.F., A.C.S., K.G.D., C.M.R., and M.D.S. wrote for broad-scale bistability. the paper. The authors declare no competing interest. alternative stable states | tropical biomes | tree species composition | This article is a PNAS Direct Submission. precipitation and seasonality | fire Published under the PNAS license. 1To whom correspondence may be addressed. Email: [email protected]. ree cover and canopy openness are commonly used to dif- This article contains supporting information online at https://www.pnas.org/lookup/suppl/ Tferentiate tropical forests and savannas, but the difference doi:10.1073/pnas.2011515117/-/DCSupplemental. between the two biomes is not just a matter of structure (1). www.pnas.org/cgi/doi/10.1073/pnas.2011515117 PNAS Latest Articles | 1of8 Downloaded by guest on September 27, 2021 Whereas tropical moist forests form a closed canopy with a used georeferenced herbarium collections (24, 25) to expand the complex vertical structure, savannas are more open, allowing floristic information spatially, to describe the complete distri- fire- and drought-adapted grasses to grow in the understory. bution of forest and savanna across Africa, and to identify the Typically, forest dominates in wetter areas, while savannas occur determinants of their present day distribution. in drier, seasonal areas (1, 2), although transitions between forest and savanna are not rigidly determined by climate (2–4). Soils Extensive Climatic Overlap of Forest and Savanna in Africa and topography can be locally and regionally important, but at Alongside the expected patterns of forest in wetter areas and intermediate rainfall (between 1,000 and 2,500 mm globally), savanna in drier and more seasonal ones (1, 2), also retrieved forest and savanna, both widespread, potentially represent al- here (Fig. 1A and SI Appendix, Fig. S1), we found an extensive ternative stable states maintained by feedbacks between tree climatic area (Fig. 1 B and C and SI Appendix, Fig. S2) within cover and disturbances—specifically fire (3, 5–7) and chronic which both forest and savanna sites are widespread, which we herbivory (8). As a result, forest and savanna tree species show term the “bistable” region. Mean annual precipitation (MAP) contrasting adaptations (9, 10), and transitions across the partly differentiates the forest, the bistable region, and the sa- forest–savanna boundary are characterized by high species vanna, although the climatic gradient used to determine and map turnover (10, 11). them is more complex and integrates precipitation and season- Despite these functional and floristic differences, most recent ality (Materials and Methods has the details of the principal efforts devoted to understanding forest–savanna transitions have component analysis [PCA] on gridded climate data).
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